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The Evolution and Diversification of


Kim Lindgren

Euphrasia stricta

Genetics and Evolution, 2hp

EMG, Umeå University.

Copyright 2010 © Kim Lindgren (

This work is licensed under a Creative Commons Attribution-No Derivative Works 3.0 Unported Licence

The angiosperms (flowering plants) are, despite their relatively recent origin, an extremely
ecologically and morphologically diverse group of plants (Soltis and Soltis, 2004). The name
angiosperm comes from the greek word angion, meaning “container” (Campbell et al., 2009;
Niklas, 1997), which refers to the seeds being hidden from sight (Niklas, 1997). The clade
consists of about 250,000-300,000 living species (Campbell et al., 2009; Niklas, 1997; Soltis
and Soltis, 2004) in about 450 families (Soltis and Soltis, 2004), that dominate the plant-life
in most terrestrial environments (Campbell et al., 2009; Qiu et al., 1999), and have done so
for about 90 million years (Qiu et al., 1999).

The origin and diversification of this clade has perplexed the scientific community for more
than 100 years, due to uninformative fossil records (Qiu et al. 1999), uncertain relationships
between living angiosperms (Kubitzki, 1993) and difficulty in identifying the earliest
angiosperms and their sister group (Qiu et al., 1999). Only recently (the last decade) have we
been able to establish the basal nodes of the angiosperm clade with a high degree of certainty
(Soltis and Soltis, 2004). This paper will present the results of recent studies in the area of
angiosperm evolution and diversification, a subject that has fascinated scientists for over a
century, something that Darwin himself called an “abominable mystery” (Campbell et al.,


Floral morphology is extremely diverse (Soltis et al. 2005), which is due to the need to spread
pollen from one plant to another (Niklas, 1997). Some flowers are very small and have water
and wind borne pollen, while some flowers are very large and use insects or even mammals
and birds to spread their pollen (Niklas, 1993). Angiosperm evolution has even yielded
flowers that mimic the mates or prey of pollinating insects; while some produce numerous
fleshy parts that are offered as food to attract pollinators (Niklas, 1997). All flowers are,
however, united by a set of synapomorphies (Soltis et al., 2005). Even though it may be
difficult to imagine, all the parts of the flower (Septals, Petals, Stamens and Carpels) are
actually highly modified leaves (Niklas, 1997). The petals and septals together form the
perianth (which means “around the flower”), that sits around the stamens and carpels (Niklas,
1997). In most species, the septals are green and scaly and serve to cover the flower as it
matures (Mossberg and Stenberg, 2003; Niklas, 1997), in others, the septals are large and
colorful, to complement or completely replace the petals as a mechanism to attract pollinators
(Niklas, 1997). The stamen bears microsporangia that produce the pollen, while the carpel
bears one or more integumented megasporangia (ovule) (Niklas, 1997). The angiosperm
ovules are surrounded with two integuments (a membrane that later becomes a seed coat),
unlike their gymnosperm relatives (see Figure 1) who only have one (Niklas, 1997).

It’s widely accepted that pollinators have affected floral evolution (Galen, 1996; Niklas,
1997). Studies on Polemonium viscosum have shown that large and colorful flowers attract
more pollinators, even though the nectar yield is at the same level or lower than in smaller
flowers (Galen, 1996).


The fossil record of angiosperms extends back to the early Cretaceous (roughly 140 million

years) (Campbell et al., 2009; Soltis and Soltis, 2004) and the group experienced an explosive
increase during this period (Campbell et al., 2009; Qiu et al., 1999). Floral diversity among
the early angiosperms was extensive, which is consistent with explosive diversification and
radiation (Soltis and Soltis, 2004). Some have hypothesized that different lineages of
angiosperms have different ancestors, however, all molecular analyzes that have been made
strongly rejects these hypotheses (Soltis et al., 2005).

Figure 1: A phylogeny showing the most probable

relationships between extant land plants, based on
molecular and morphological data. The darker lines
and text show the angiosperm clad (Adapted from
Soltis et al., 2005 and Campbell et al., 2009).

The closest living relatives to the angiosperm clad are the gymnosperms (Campbell et al.,
2009; Soltis et al., 2005) and morphological and molecular evidence suggests that the
gymnosperms diverged from the ancestor of the angiosperms roughly 300 million years ago
(Campbell et al., 2009). However the fruits and flowers that are unique to the angiosperms
distinguish them greatly from their relatives, which is one of the reasons why the origin of the
angiosperms has puzzled so many for so long (Campbell et al., 2009), although, some modern
fossil findings of extinct gymnospem species show reproductive structures that are similar to
those found in the angiosperms (Niklas, 1997). It is also possible that the angiosperms are in
fact more closely related to a group of extinct seed plants, known as Bennettitales, which had
a flower-like structure (Campbell et al., 2009). Figure 1 shows the relationships between
different groups of extant angiosperms, as well as their relation to other groups of extant land

The most probable reason why they rose to their dominant position so quickly, are some of the
important reproductive features that other seed plants lack, such as: protected ovules, stigmas,
double fertilization and accelerated gametophyte development (Furness and Rudall, 2003;
Kubitzki, 1993).


This section will describe the three most basal nodes of the angiosperm phylogeny. These
three lineages – Amborella, Nymphaeaceae and Austrobaileyales (Campbell et al., 2009;
Soltis et al., 2005) – are comprised of only about 100 extant species (Campbell et al., 2009),
the earliest consisting of only a single species (Campbell et al., 2009; Soltis and Soltis, 2004;

Soltis et al., 2005; Williams, 2009). The remaining angiosperm clades (Magnoliids,
Chloranthaceae, Ceratophyllaceae, Monocotyledons and Eudicots, see Figure 1), are referred
to as core angiosperms (Magallón and Castillo, 2009). Common to all basal angiosperms are
pollen with only a single aperture, a trait shared with the gymnosperms (Furness and Rudall,
2003; Soltis et al., 2005) and the core angiosperms, except the eudicots (Furness and Rudall,

4.1 Amborella trichopoda

The current consensus is that Amborella trichopoda (The only currently living member of the
Amborellaceae family), a small understory evergreen tree endemic to New Caledonia
(Campbell et al., 2009; Soltis and Soltis, 2004; Soltis et al., 2005; Williams, 2009), which is
both wind and insect pollinated (Williams, 2009), should be placed as the sister to all other
living angiosperms (Campbell et al., 2009; Soltis and Soltis, 2004). Although, some studies
suggest that A. trichopoda should occupy this position along with Nymphaeaceae (described
below) (Soltis and Soltis 2004; Soltis et al., 2005).

A. trichopoda have features that may be ancestral to the angiosperms, such as carpels that are
only closed by secretion, rather than by fused tissue (Soltis et al., 2005), and unlike all other
known extant angiosperms, the xylem of A. trichopoda lack vessel elements (Soltis et al.,
2005). The pollen is tectate (Philipson, 1993), and the aperture is less pronounced than in the
core angiosperms (Philipson, 1993; Soltis et al. 2005). The process in which A. trichopoda
produces eggs is also very similar to the equivalent process in gymnosperms (Campbell et al.,

4.2 Nymphaeaceae

The position of Nymphaeaceae as one of the earliest diverging clad of angiosperms is

supported by almost all genetic studies (Soltis and Soltis, 2004; Soltis et al., 2005) and it was
probably predated only by Amborella (Campbell et al., 2009). However, even though the
group diverged early from other angiosperms, most extant members of the Nymphaeaceae
family have a relatively recent origin (Soltis et al. 2005). They have a worldwide distribution
and are fully aquatic, which is consistent with the age of the clad (Soltis and Soltis, 2004).
Their flowers are extensively diverse (Soltis and Soltis, 2004), which is also consistent with
their age, and is most probably due to the suitability of different pollinators and different
methods of pollination in different habitats (Soltis and Soltis, 2004). The pollen is, like the
flowers, very diverse, some with more archaic features, such as the pollen being tectate and
having a smooth surface, as well as the aperture of the pollen being less pronounced (like in
A. trichopoda) than in the core angiosperms (Schneider and Williamson, 1993).

4.3 Austrobaileyales

This small group of large lianas (Endress, 1993) that can be found in Australia and Asia
follows Amborella and Nympaeaceae as the successive sister to the rest of the angiosperms
(Soltis et al. 2005). Unlike A. trichopoda and Nymphaeaceae the aperture on the pollen of
Austrobaileyales is more similar to that of the core angiosperms (Endress, 1993), however the
pollen still have some very primitive characters (Endress, 1993), such as the outer wall of the
pollen being tectate-perforate and supported by a columellate structure (Endress, 1993).


For a long time (up to the late 1990s), the angiosperms where divided into two groups: the
monocots and the dicots (Kubitzki, 1993, Soltis et al., 2005). More recent studies on
angiosperm genetics have however shown that, even though the monocots do in fact form a
monophyletic group, the dicots do not (Soltis et al., 2005). Most of the angiosperms
traditionally seen as dicots are now found in the eudicot clad (true dicots), however; A.
trichopoda, Nymphaeaceae, Austrobaileyales, Ceratophyllaceae, Chloranthaceae and the
Magnoliids (see Figure 1), also used to belong to the dicot clad (Soltis et al., 2005).

5.1 Monocotyledons (monocots)

The monocots where first identified in 1703 (Soltis et al., 2005), and is a group strongly
supported by both molecular studies and a set of distinct synapomorphies (Soltis et al., 2005).
These include having only a single seed leaf (which gave the group its name), but also by the
leaf veins growing parallel to each other, the xylem is scattered and they have fibrouse roots
that lack a main structure (Campbell et al., 2009; Soltis et al., 2005). Like the basal
angiosperms, monocot pollen has only one aperture (Campbell et al., 2009; Furness and
Rudall, 2003; Soltis et al., 2005). Even though monophyly for the monocot clad is strongly
supported, their relationship to the other angiosperms is unclear (Soltis et al., 2005).

5.2 Eudicots

This large clad contains most of the angiosperm species (roughly 75%), and is very
morphologically diverse (Furness and Rudall, 2003; Soltis et al., 2005). Besides strong
molecular evidence for the clads monophyly, only a single synapomorphie unites the eudicots:
pollen with three or more apertures (Campbell et al., 2009; Soltis et al., 2005), which is
unique to the eudicots (Furness and Rudall, 2003; Soltis et al., 2005). Even though they are
not synapomorphies, the xylem in the eudicots is generally arranged as a ring, as well as the
leaves normaly having net-like veins, and the roots being more structured than in monocots
(Campbell et al., 2009).

The feature of having three or more apertures is probably the main reason for eudicot success
(Furness and Rudall, 2003), since they increase the chances of germination and thus have the
potential to increace the rate of fertilization (Furness and Rudall, 2003).


Angiosperms have an advanced reproductive system, which probably is the reason why they
so quickly rose to their dominant position in the plant kingdom (Kubitzki, 1993), as well as
the reason for the great morphological and ecological diversity in the clad. Successive
adaptations to pollen and flowers that attract pollinators have increased reproductive success,
and enabled the angiosperms to spread their genetic material over greater distances. Even
though these features are costly (Galen, 1996), they give the angiosperms an advantage over
other seed plants. Further adaptations in pollen and flower morphology have greater increased
the dominance of the angiosperm clad, by increasing the attraction of pollinators and
increasing the rate of fertilization.

The subject is very interesting, however there seems to be a general lack of books and papers

that give a clear timeline on the evolutionary stages of the flower and its components. The
literature seems to generally focus on the earliest angiosperms and their relation to other
groups, or on describing modern flowers, which, although important, leaves me wanting
more. However, I suppose that is the reason why I chose to become a biologist.

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2009. Biology. International 8th Edition. Pearson Benjamin Cummings. San Francisco. USA.

Endress P. K. 1993. Austrobaileyales. In: The Families and Genera of Vascular Plants. Kubitzki K.,
Rohwer J.G., and Bittrich V. (Eds.). Springer-Verlag. Berlin. Germany.

Furness C. A. and Rudall P. J. 2004. Pollen aperture evolution – a crucial factor for eudicot success?.
Trends in Plant Science. 9: 1360-1385

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plastid and nuclear genomes. Nature 402: 404-407

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origins of the angiosperm progamic phase. American Journal of Botany 96: 144–165.